The First Humans in Europe

Hominins, Neanderthals and the humans before the Aurignacians

Where to begin with the topic of who were the first humans in Europe? Even to ask the question is to invite the response, ‘who counts as human?’, and there’s some truth here, where do we draw the line between the different stages of human evolution? I’ll make it simple, lets start at the very beginning. Who inhabited Europe and when? Traditionally the sequence has usually run something like: unknown humans > Neanderthals > Aurignacian Homo sapiens. But we know now that the Aurignacians were not the first modern humans to set foot in Europe, and the growing body of evidence for multiple incursions deserves looking at in detail. So let’s jump in, who was the first human ancestor on the continent?

Apes, Monkeys & Migrations

In each great region of the world the living mammals are closely related to the extinct species of the same region. It is therefore probable that Africa was formerly inhabited by extinct apes closely allied to the gorilla and chimpanzee; and as these two species are now man's nearest allies, it is somewhat more probable that our early progenitors lived on the African continent than elsewhere. But it is useless to speculate on this subject, for an ape nearly as large as a man, namely the Dryopithecus of Lartet, which was closely allied to the anthropomorphous Hylobates, existed in Europe during the Upper Miocene period; and since so remote a period the earth has certainly undergone many great revolutions, and there has been ample time for migration on the largest scale

-Darwin, 1871, The Descent of Man

Darwin’s discussion of human origins notes a mystery which is still with us today, where did the earliest Great Apes originate? Our nearest relatives, the Hominidae family group, contains the chimpanzee, the orangutan, the gorilla, the bonobo and every type of human. Classifying the various human sub-species is a challenge for this century, and there’s new material turning up by the day. At first glance it would seem logical that the Great Apes also evolved in Africa, but a growing body of evidence points towards Europe as the cradle of the family.

The Miocene period is where we begin, somewhere between 23 and 5 million years ago. This epoch saw Africa join Eurasia; the Mediterranean evaporate and refill again, and apes evolve into their modern forms. By the end of the timeline we see the ancestors of chimps and humans finally split, opening the evolutionary path for modern humans to develop and diverge. The earliest known ape species in Eurasia is the Griphopithecus, a thick-enameled creature discovered in Germany, and dated to around 17 million years ago. He had arrived with many others of his kind from Africa, such as the Kenyapithecus, Equatorius and Nacholapithecus, all species with thick enamel, spread out from Kenya to Turkey to Germany. After another 5 million years the split between the orangutans and the rest of the apes becomes visible, with the former moving towards Southeast Asia. In Europe this new species of ape is collectively called the Dryopithecines - fruit-eating quadrupeds with a stiff back, thinner enamel, a broad thorax and powerful grasp. They were something like a modern ape, a vital part of the transition towards later hominins.

The earliest hominines are European and there are no fossil hominines in Africa for at least two million years after they first appear in Spain and France…

The best-known samples of late Miocene European hominines are about 10 to 9.5 Ma in age (Rudabánya, Can Llobateres, Ravin de la Pluie). By this time hominines had diversified substantially and in some ways mimic the diversity that existed among Pliocene hominins. Ouranopithecus was substantially larger, with much more robust jaws, thickly enameled teeth, and many other adaptations to a diet that must have emphasized the consumption of hard or tough food objects, like australopithecines. Rudapithecus and Hispanopithecus were highly suspensory, arboreal soft fruit frugivores, with cheek teeth closely resembling those of living chimpanzees. The brain of Rudapithecus is known to have been as large as that of modern chimpanzees, and it is possible that this fossil ape had cognitive capabilities similar to those of chimpanzees

-Miocene Hominids and the Origins of the African Apes and Humans, Begun, 2010

What seems clear, after all this time, is that Darwin’s suspicions were correct. Although the later ancestors of modern humans emerged in Africa, their forebears in turn had evolved in Europe, before leaving as the climate began to dry out.

A speculative evolutionary tree for the Great Apes. Paper here.

Indeed, there is another possibility, which is that the large range, diversity, and density of Miocene hominines in Europe point to a European origin, with a subsequent dispersal into Africa in the late Miocene. There is strong evidence for the latter hypothesis in data from the morphology, geochronology, and paleobiogeography of European hominines and other land mammals.8, 23

Hominines occurred in Europe at least two million years before they appeared in Africa.17

-European Miocene Hominids and the Origin of the African Ape and Human Clade, Begun et al, 2012

The number of features shared between Dryopiths and later hominins is quite large, including facial, nasal, sinoid and skull modifications and rearrangements. Their effect on later developments leading up to the final split between ancestral chimps and humans seems to have been profound.

The Great Apes were however, forced out of Europe, and many went extinct. What they brought back to Africa with them, along with their new physiology, was a more flexible and adaptive set of behavioral strategies, to manage the increased seasonality and ecological change. The next time their descendants set foot in Europe, the story would be very different.

Back-to-Europe: Homo antecessor and their kin

The first hints of something happening in Europe come from a controversial set of fossil fragments and teeth found in Greece and Bulgaria. Dubbed Graecopithecus freybergi, the remains were the centre of an academic controversy in 2017, when a paper published in PLOS One tentatively argued that Graecopithecus could be the first example of the hominin family after the break with ancestral chimpanzees. Dated to 7.2 million years ago, the media declared that Europe and not Africa was the birthplace of the human species. Other academics have responded, ultimately leading to both camps accepting that we don’t have the evidence yet to support Graecopithecus’s inclusion into the hominin group.

The first accepted appearance of the newly evolved Homo species in Europe was recently discovered in Spain. The archaeological site of Sima de los Heusos, Atapuerca, is amongst the important in the world, having yielded tools and bones of numerous human species, including Neanderthals. The genetic and skeletal data from the site has been invaluable in building family relationships between different human species. The oldest to date are between 1.4 and 1.2 million years old, which include teeth, a finger bone and two jawbones. They could belong to either Homo erectus or antecessor. The consensus leans towards them being Homo antecessor, which makes sense, since this sub-species of human has only been found in Spain.

Antecessor, or ‘pioneer man’, is one of the many mysterious branches of the human tree that we don’t fully understand. They seem to be an offshoot of the species that would evolve into Homo heidelbergensis, the common ancestor to both modern humans and Neanderthals. What is particularly eerie though is that antecessor appears to have developed very similar skeletal characteristics as modern humans, including a gracile, flatter face and similar spinal and shoulder anatomy. Almost like a dry-run for modern humans. However, the fate for hominids is never usually good, and antecessor appears to have gone extinct. Their legacy seems to have been a voracious taste for hunting down juvenile members of their own kind and cannibalising them.

This period of time, between the evolution of the Homo genus and the more familiar later species has always been an awkward muddle. Numerous species and sub-species have been named, classified, re-classified and dropped, and the debates over whether a single tooth represent a new hominin or an existing one are dull. So we shall proceed with the most straightforward of narratives as I see it, but you may disagree. Around 1.5 million years ago then, we have an offshoot species living in Spain, likely the first to arrive back from Africa, but they don’t contribute to any later surviving population. Instead we see the advance and much wider spread of a different human species - Homo heidelbergensis. This thickly-browed, fire-using species has left evidence in the form of skeletons, artefacts and even footprints as far north as Britain. From what limited evidence we have, they must have inherited the Acheulean handaxe tradition from their Homo erectus forebears and made limited forays into Europe as the weather permitted. The dates for this colonisation are all over the place, but after about 800,000 years ago, we begin to see the coalescence of adaptive behaviours which allowed humans to live in much colder climates. The use of fire, hafted weapons, handaxes and a diverse set of strategies for hunting different animals and foraging for plant foods all contributed to the later appearance of Europe’s longest lived inhabitants - the Neanderthals.

A Neanderthal World

In comparison to the feeble evidence we have for Europe’s humans over the hundreds of thousands of years that antecessor and heidelbergensis lived and died, evidence for Neanderthals seems plentiful. Exactly when they split from the rest of the human family is not clear, somewhere between 400 and 800 thousand years ago, but they flourished in Europe until around 40,000 years ago - meaning they may have inhabited the continent for ten times longer than modern Homo sapiens. Their exploits have become better known in recent decades, rising from their poor early PR as primitive savages to being respected as an inventive, creative species - capable of sailing, making fire, distilling tar, making art, hunting huge animals, producing clothes, potentially venerating eagles and identifying medicinal plants. We also know a great deal more about their genetics, and when and where they interbred with modern humans and Denisovans.

The details of how Neanderthals lived, their hunting strategies, tool production, domestic settings and so on have been well documented in many recent books and articles. What is often missing from academic archaeology is any kind of meta-narrative, how populations formed, moved and disappeared. From this vantage Neanderthals are still somewhat mysterious, and they seem to behave very differently in their wider social relations than modern humans. Their evolution from Homo heidelbergensis seems the most likely origin story, with small groups being isolated as successive waves of glaciation hit Europe, over time forming the distinctive Neanderthal features in at least Western Europe, before they went east, all the way to Siberia. By 120,000 years ago there were at least two deeply divergent Neanderthal populations, generally defined by ‘western’ and ‘eastern’ titles.

The deepest divergence among Neanderthal genomes sequenced to date is found between eastern and western Eurasian Neanderthal populations represented by the ~120 ka Altai Neanderthal from Denisova Cave7 and the >44 ka Vindija 33.19 individual from Croatia. Genomic data of all other available Neanderthal remains, the earliest in western Europe being ~120 ka (Scladina and Hohlenstein-Stadel (HST)), while the latest being ~40 ka, suggest genetic continuity in western Eurasia for ~80 ka. Recent results obtained from sedimentary DNA suggest that the genetic landscape was significantly altered by expansions of Neanderthal populations ~105 ka. This gave rise to lineages in western Europe represented by samples from Central Europe (Vindija), the Caucasus (Mezmaiskaya Cave), and Siberia (Chagyrskaya cave 8), the latter likely replacing the earlier Altai-like population. The genomes of late (<50 ka) European Neanderthals, including an individual from the Caucasus (Mezmaiskaya 2), were all found to be more similar to Vindija than to other known lineages, indicating further population turnover towards the last stages of Neanderthal history in the Caucasus or western Europe.

However, even within these broad lineages and turnovers, Neanderthals seem to differ from modern humans in their extremely low inter-group mixing and higher instances of inbreeding. A recent discovery of a well-preserved Neanderthal male from southern France, nicknamed ‘Thorin’, revealed that Europe possessed a number of small, isolated groups, which kept to themselves for millennia.

Our results nevertheless suggest a minimum of two distinct Neanderthal lineages present in Europe during the late Neanderthal period. In the absence of any detectable gene flow between Thorin and other Neanderthal lineages after its divergence, we conclude that Thorin represents a lineage that has stayed isolated for ~50 ka… Our results thus also shed light onto the social organization of Neanderthals, suggesting that small isolated populations with limited, and potentially without, inter-group exchange as a possibly more general feature of Neanderthal social structure.

Best fitting model for Thorin and his relationships to other Neanderthals and Denisovans - Slimak et al 2023

The divergence of Thorin’s lineage also matches another discovery from northern Spain, where an almost total Neanderthal population replacement occurred around 100,000 years ago. Clearly over a very long time span, Neanderthals were also separating, remaining separate and moving into one another’s territory. Evidently they also preferred to live and mate amongst their own groups. One has to imagine that isolation for tens of thousands of millennia would produce huge language differences, assuming language development worked in the same way for Neanderthals as modern humans.

Waves of Humans

Dating the entrance of Homo sapiens into Europe has always been a topic of great interest amongst archaeologists, especially since examples dating back around 180,000 years have been found at Misliya Cave in Israel. Debates continue to be had over the identity of a pair of skulls found at Apidima Cave in Greece, one of which has been suggested to be an early Homo sapien, but could equally be Homo erectus with early Neanderthal features. Given that these date back around 210,000 years, we are looking at potentially multiple waves of modern humans either within Europe or on her borders.

If Thorin was an exceptional find for archaeology, his burial place might enter the top ten sites for all of prehistory. The cave of Grotte Mandrin is set to rewrite the human story, and provides the best and most detailed sequence of events surrounding the first attempts of modern humans to colonise Europe. The limestone cave system is perched high above the Rhône valley, providing superlative views of the landscape, access to river systems and good quality flint. The cave provides an almost perfect set of stratigraphic layers which document not only the lives of its resident Neanderthals from 100kya to their extinction 42kya, but also astonishing glimpses into the earliest Homo sapien visits into the continent. Around 120,000 flint tools and animal bones have been painstakingly recovered, along with millions of micro flint chips, and many human remains. As I’ve written before, deep prehistory tends to be defined by types of stone tool industry, in this case the Mousterian being the style associated with Neanderthal sites and skeletal remains.

The final series of phases from 54kya to 42kya seems to run thus:

1. Mousterian style stone tools - Neanderthal

2. Short transition period dubbed ‘Neronian’ - modern humans

3. Two post-Neronian phases with Mousterian tools - Neanderthal re-occupation

4. First Upper Palaeolithic layer - ‘proto Aurignacian’ modern humans

The distinguishing features here are the styles of stone tools. To the layman, this might sound a bit vague, but the contrast between the Mousterian tools, which are large flakes, and the Neronian is huge. We can look not only at the final form of the tools, which are nothing alike, but also the manufacturing methods required. The Neronian toolkit is dominated by excessively small micro or nano points, which have no precedent in the hundreds of millennia in Europe before then. These tiny crafted weapons point in one direction - the first use of the bow-and-arrow.

Neronian micro points found at Grotte Mandrin

Not only do comparative studies of these points against experimental flint arrowheads further support this hypothesis, but they hint that the Neronian arrows may have been tipped with poison.

Incredibly the researchers working on the cave have been able to use trapped layers of soot in the interior to date the sequence of occupations, akin to dating a tree by its rings. In an astonishing set of results, archaeologists were able to determine that no more than one or two years passed between the Neanderthals and the first human Neronians. These were no mere scouts, a child’s tooth was discovered in the Neronian layer, and they maintained a presence for around forty years before disappearing, the cave falling back into the hands of the Rhône valley Neanderthals.

What happened next is just as remarkable. The controversial tool industry known as the Châtelperronian has been argued to be either a Neanderthal innovation based on contact with modern humans, or an early intrusion of humans into Neanderthal Europe. Recent arguments for the Châtelperronian sites in Franco-Iberia as a second wave of sapiens has to be considered alongside the reoccupation of the Rhône:

If the Châtelperronian effectively corresponds to a second migratory phase by H. sapiens, and originated from the same Levantine cultural substrate, the absence of chronological and geographical overlap between phase I (IUP / Neronian) and phase II (NEA / Châtelperronian) is all the more remarkable, as the territorial expansion of this phase II affected large territories- Atlantic, continental, and Mediterranean- which remain quite geographically disjointed. Over this same period, the Rhône valley was occupied by Neandertal groups that carried the Post-Neronian II traditions [1]. Could it be that in the same geographical space that saw the first migrations of H. sapiens into Europe, Neanderthal groups no longer allowed access to their previous territory? This would be remarkable, since the Post-Neronian I and Post-Neronian II, which mark a return of Neandertal populations to a large territory around Mandrin, also indicate a persistence of Neandertal populations in one of the main migratory arteries of Western Europe [1]. This could well indicate a refusal or a resistance from the aboriginal populations against a return of H. sapiens at the very moment when, according to this hypothesis, these latter populations would manifest their first real colonization by way of settlements, not only numerous, but also over vast territories across Western Europe.

What we have to postulate then is that modern humans had established a strong maritime presence along the Mediterranean shorelines, and were finding major river systems blocked off by Neanderthals. They were forced to instead move up through northern Iberia and the Balkans.

The final proto Aurignacian expansions, circa 42kya, is visible within Grotte Mandrin. Again soot deposits reveal the overlap to be a few years at most, perhaps shorter. This wave of colonisation was to be the last time Neanderthals lived in the valley, their ancestral home taken over by the new arrivals. It is entirely possible that these last Neanderthals were using small point weaponry, having inherited the tradition from a long forgotten Neronian population over ten thousand years prior.

Current model of pre-Aurignacian human expansion into Europe - from Slimak 2023

The Last Neanderthals

We still have a long way to go to understand these pre-Aurignacian human groups. One of the only sets of human remains from this period, dated to 45,000 years ago, is the Zlatý kůň skull from the Czech Republic. Her genome showed similar levels of Neanderthal introgression, but the sequences were much longer, and she did not pass on her ancestry to the later Aurignacians and Gravettians. As for the Neronians and Châtelperronians, we are currently blind. Hopefully DNA can be extracted from those found at Grotte Mandrin, which would give us much more information about who was arriving on the shores of Europe over 50kya.

The Neanderthals themselves suffered not only the colonisation of their lands by the sapiens, but were also hit with a major climatic downturn around 42,000 years ago. The abandonment of inner Iberia, coincided with the Protoaurignacian ascendency. The reasons for the Neanderthal’s demise are beyond what I can write here, but likely a combination of genetic fitness, social flexibility and adaptability, higher birthrates, use of dogs and multiple niche exploitation all counted. Aggression and violence seem a given, over the millennia of interactions, and since the Neanderthals seemed to have avoided each other, they likely shunned the newcomers as well for the most part.

As to their survival, it is possible that some clung on past 40,000 years ago. A number of disputed sites exist within Spain, and a controversial skeleton of a young boy found at Lagar Velho 1 in Portugal hints at some longevity. Dated to 24,500ya, the boy reportedly shows facial features and anatomical structures consistent with being a Neanderthal hybrid, but other researchers have argued this is a preservation issue. At least one paper argues for an Iberian holdout up until 28kya, and the Croatian Vindija Cave Neanderthals have been redated to around 32kya. Probably the best site comes from the Polar Urals, at a site called Byzovaya, where Mousterian tools were tentatively dated to around 33,000 years ago. The site contains butchered mammoths and reindeer, with contemporary Upper Palaeolithic stone tool sites close by. Along with the Russian Mezmaiskaya Cave Neanderthal bone dates of 35kya, it is looking more and more like Neanderthals retreated into two refugia - one in southernmost Iberia and one towards the Russian Arctic. Since no Mousterian tools have been associated with modern humans, this doesn’t seem a far-fetched hypothesis.

Conclusions

So we come to the end of this long tale, one which has seen various stages of human evolution take place within Europe. The time scales are of course staggering, and we can’t really imagine what 50k years of isolation looks like, let alone 500k or a million. What we can see are broad trends, supplemented by amazing snapshots of moments in time. The slow domination by hominins who possess greater adaptability, both biological and cultural, seems to be the story of Europe’s various human branches. The latest developments in building something of a narrative to understand the arrival of H. sapiens has to be amongst the most exciting topics in archaeology today - to be able to see in soot depositions just a few years in a person’s life is mind-blowing, and being able to deduce from pieces of stone that the first intruders carried a new and deadly projectile weapon is fodder for the imagination. The story of Europe’s Neanderthals, melting away as the sapien trail is blazed, is also poignant. Surviving either on the extremities of Iberia, perhaps looking to Africa to escape, or butchering mammoth in the chill dark of the long polar night, their backs turned to their sapien neighbours just over the hill. Hopefully we find more of these stories to tell, the deep time of Europe’s first humans.